Susan K. Skagen

NEST SURVIVAL ESTIMATION: A REVIEW OF ALTERNATIVES TO THE MAYFIELD ESTIMATOR

Abstract Reliable estimates of nest survival are essential for assessing strategies for avian conservation. We review the history of modifications and alternatives for estimating nest survival, with a focus on four techniques: apparent nest success, the Mayfield estimator, the Stanley method, and program MARK. The widely used Mayfield method avoids the known positive bias inherent in apparent nest success by estimating daily survival rates using the number of exposure days, eliminating the need to monitor nests from initiation. Concerns that some of Mayfields assumptions were restrictive stimulated the development of new techniques. Stanleys method allows for calculation of stage-specific daily survival rates when transition and failure dates are unknown, and eliminates Mayfields assumption that failure occurred midway through the nest-check interval. Program MARK obviates Mayfields assumption of constant daily survival within nesting stages and evaluates variation in nest survival as a function of biologically relevant factors. These innovative methods facilitate the evaluation of nest survival using an information-theoretic approach. We illustrate use of these methods with Lark Bunting (Calamospiza melanocorys) nest data from the Pawnee National Grassland, Colorado. Nest survival estimates calculated using Mayfield, Stanley, and MARK methods were similar, but apparent nest success estimates ranged 1– 24% greater than the other estimates. MARK analysis revealed that survival of Lark Bunting nests differed between site–year groups, declined with both nest age and time in season, but did not vary with weather parameters. We encourage researchers to use these approaches to gain reliable and meaningful nest survival estimates. Estimación de la Supervivencia de Nidos: Una Revisión de las Alternativas del Estimador Mayfield Resumen. Es esencial contar con estimaciones confiables de la supervivencia de nidos para evaluar las estrategias de conservación de las aves. Revisamos la historia de modificaciones y las alternativas para estimar la supervivencia de nidos, enfocándonos en cuatro técnicas: éxito aparente del nido, el estimador de Mayfield, el método de Stanley y el programa MARK. El método de Mayfield, ampliamente usado, evita el conocido sesgo positivo inherente al éxito aparente del nido mediante la estimación de tasas de supervivencia diaria usando el número de días de exposición, eliminando así la necesidad de monitorear los nidos desde el inicio. Las preocupaciones de que algunos de los supuestos del método de Mayfield son restrictivos estimularon el desarrollo de nuevas técnicas. El método de Stanley permite el cálculo de tasas de supervivencia diarias específicas para cada etapa cuando las fechas de transición y fracaso son desconocidas, y elimina el supuesto del método de Mayfield que sostiene que el fracaso ocurre en el medio del intervalo de monitoreo del nido. El programa MARK elimina el supuesto del método de Mayfield sobre supervivencia diaria constante dentro de las etapas de nidificación y evalúa la variación en la supervivencia de nidos como función de factores biológicamente relevantes. Estos métodos innovadores facilitan la evaluación de la supervivencia de nidos usando un enfoque teórico-informativo. Ilustramos el uso de estos métodos con datos de nidos de Calamospiza melanocorys provenientes de Pawnee National Grassland, Colorado. Las estimaciones de supervivencia de los nidos calculadas usando los métodos de Mayfield, Stanley y MARK fueron similares, pero las estimaciones del éxito aparente de los nidos fueron entre 1–24% mayores que las otras estimaciones. Los análisis con MARK revelaron que la supervivencia de los nidos de C. melanocorys difirió entre grupos de sitio-año, disminuyó con la edad del nido y el tiempo de la estación, pero no varió con parámetros climáticos. Estimulamos a los investigadores a usar estos enfoques para obtener estimaciones de supervivencia de nidos confiables y válidas.

MODELING POST-FLEDGING SURVIVAL OF LARK BUNTINGS IN RESPONSE TO ECOLOGICAL AND BIOLOGICAL FACTORS

We evaluated the influences of several ecological, biological, and methodological factors on post-fledging survival of a shortgrass prairie bird, the Lark Bunting (Calamospiza melanocorys). We estimated daily post-fledging survival (n = 206, 82 broods) using radiotelemetry and color bands to track fledglings. Daily survival probabilities were best explained by drought intensity, time in season (quadratic trend), ages < or = 3 d post-fledging, and rank given drought intensity. Drought intensity had a strong negative effect on survival. Rank was an important predictor of fledgling survival only during the severe drought of 2002 when the smallest fledglings had lower survival. Recently fledged young (ages < or = 3 d post-fledging) undergoing the transition from nest to surrounding habitat experienced markedly lower survival, demonstrating the vulnerable nature of this time period. Survival was greater in mid and late season than early season, corresponding to our assumptions of food availability. Neither mark type nor sex of attending parent influenced survival. The model-averaged product of the 22-d survival calculated using mean rank and median value of time in season was 0.360 +/- 0.08 in 2001 and 0.276 +/- 0.08 in 2002. Survival estimates that account for age, condition of young, ecological conditions, and other factors are important for parameterization of realistic population models. Biologists using population growth models to elucidate mechanisms of population declines should attempt to estimate species-specific of post-fledging survival rather than use generalized estimates.

Human disturbance of an avian scavenging guild

In order to investigate the effects of human activities on relationships within foraging guilds, we examined inacanus dynamics of eagles, crows, and gulls scavenging on spawned salmon in the Pacific Northwest. We examined several hypotheses that postulate the asymmetric foraging relationships of the three guild members and that reveal the influence of competition and facilitation in these relationships. Spatial and temporal patterns of resource use by the three primary guild members varied with the presence and absence of human activity at experimental feeding stations. At control (undisturbed) stations, eagles preferred to feed >100 m from vegetative cover, whereas gulls fed <50 m from cover. At experimental (disturbed) stations, eagles rarely fed, and feeding activity by gulls increased at both near and far stations. Crows often fed on alternate food sources in fields adjacent to the river, especially when salmon carcasses were scarce, whereas eagles and gulls rarely did so. We also examined if and how the behavior of single guild members changes in the presence or absence of other guild members. In the absence of eagles, gulls and crows preferred stations far from cover, numbers of both increased at feeding stations, birds were distributed nearer to carcasses, and they fed more. We emphasize that guild theory lends important insights to our understanding of the effects of human disturbance on wildlife communities.

Residency patterns of migrating sandpipers at a midcontinental stopover

Arctic-nesting shorebirds require several refueling stops during their long migrations between breeding grounds and Central and South American wintering areas. The protection of stopover habitats for transcontinental migrants depends on whether birds fly long distances between a few select sites or fly short distances and stop at several wetlands. Although the Great Plains historically provided a vast array of wetlands for use by migrants, wetland loss and conversion have reduced the availability of stopover sites in recent decades. In this study, we examined (1) residency periods, (2) fat dynamics, and (3) migration chronology of two shorebird species, the Semipalmated Sandpiper (Calidris pusilla) and Whiterumped Sandpiper (C. fuscicollis) at Quivira National Wildlife Refuge (NWR), Kansas. Semipalmated Sandpipers had prolonged periods of species residency with overlapping arrivals and departures. Individual residency periods were highly variable and were unrelated to lipid reserves upon arrival. In contrast, White-rumped Sandpipers arrived and departed more synchronously. Birds that arrived in poor condition stayed longer than those with more body fat in 1991, but not in 1992. Wind direction did not influence patterns of departures of either species. We hypothesize that Semipalmated Sandpipers are ecologically eurytopic when migrating across the Great Plains in the spring. Highly variable patterns in arrival, residency, and lipid levels indicate that spring migration of this species is relaxed and opportunistic. White-rumped Sandpipers showed a pattern of reduced flexibility. Flight range estimates suggest that most birds require intermediate stopovers before reaching the breeding grounds. Interior wetlands appear to function as migration stopovers rather than staging areas for shorebirds.

MOVEMENTS AND SURVIVAL OF LARK BUNTING FLEDGLINGS

Abstract We quantified post-fledging pre-independence behavior and survival in Lark Buntings (Calamospiza melanocorys) using radio-telemetry. Brood division was recorded in six broods and was maintained throughout the observed fledgling care period. Chicks were capable of short flights (up to 25 m) by fledgling day 6 and longer flights (to 100 m) by fledgling day 13. During the first three weeks after fledging, juveniles moved as far as 800 m from nests. Nine of 23 (39%) monitored fledglings died within 15 days of fledging, primarily due to predation by raptors. Daily survival rates were 0.953 ± 0.019 for fledgling days 0–9, 0.955 ± 0.038 for fledgling days 10–20, and 0.953 ± 0.015 for fledgling days 0–20. The probability of surviving fledgling days 0–20 was 0.367. More quantification of juvenile survival is clearly needed to understand the role of post-fledging mortality in source-sink dynamics. Los Movimientos y Supervivencia de los Volantones de Calamospiza melanocorys Resumen. Cuantificamos la conducta y la supervivencia de volantones de Calamospiza melanocorys antes de independizarse de sus padres usando telemetría de radio. La división de la nidada se registró en seis nidadas y se mantuvo a través del período del cuidado de los volantones. Tras seis días de haber abandonado el nido, los polluelos eran capaces de realizar vuelos cortos (de hasta 25 m) y para el día trece ya realizaban vuelos más largos (a 100 m). Durante las primeras tres semanas después de salir del nido, los juveniles se movieron hasta 800 m de los nidos. Nueve de 23 (39%) volantones se murieron en los primeros 15 días fuera del nido, principalmente debido a depredación por aves rapaces. Las tasas diarias de supervivencia fueron de 0.953 ± 0.019 para los días 0 a 9, 0.955 ± 0.038 para los días 10 a 20, y 0.954 ± 0.015 para días 0 a 20. La probabilidad de sobrevivir entre los días 0 y 20 fue de 0.367. Se necesita más cuantificacion de la sobrevivencia en la etapa juvenil para entender el papel de la mortalidad tras la salida del nido en la dinámica de fuente-sumideros.

Weather effects on avian breeding performance and implications of climate change.

The influence of recent climate change on the worlds biota has manifested broadly, resulting in latitudinal range shifts, advancing dates of arrival of migrants and onset of breeding, and altered community relationships. Climate change elevates conservation concerns worldwide because it will likely exacerbate a broad range of identified threats to animal populations. In the past few decades, grassland birds have declined faster than other North American avifauna, largely due to habitat threats such as the intensification of agriculture. We examine the effects of local climatic variations on the breeding performance of a bird endemic to the shortgrass prairie, the Lark Bunting (Calamospiza melanocorys) and discuss the implications of our findings relative to future climate predictions. Clutch size, nest survival, and productivity all positively covaried with seasonal precipitation; yet relatively intense daily precipitation events temporarily depressed daily survival of nests. Nest survival was positively related to average temperatures during the breeding season. Declining summer precipitation may reduce the likelihood that Lark Buntings can maintain stable breeding populations in eastern Colorado although average temperature increases of up to 3 degrees C (within the range of this study) may ameliorate declines in survival expected with drier conditions. Historic climate variability in the Great Plains selects for a degree of vagility and opportunism rather than strong site fidelity and specific adaptation to local environments. These traits may lead to northerly shifts in distribution if climatic and habitat conditions become less favorable in the drying southern regions of the Great Plains. Distributional shifts in Lark Buntings could be constrained by future changes in land use, agricultural practices, or vegetative communities that result in further loss of shortgrass prairie habitats.

Estimation of lipids and lean mass of migrating sandpipers

Estimation of lean mass and lipid levels in birds involves the derivation of predictive equations that relate morphological measurements and, more recently, total body electrical conductivity (TOBEC) indices to known lean and lipid masses. Using cross-validation techniques, we evaluated the ability of several published and new predictive equations to estimate lean and lipid mass of Semipalmated Sandpipers (Calidris pusilla) and White-rumped Sandpipers (C. fuscicollis). We also tested ideas of Morton et al. (1991), who stated that current statistical approaches to TOBEC methodology misrepresent precision in estimating body fat. Three published interspecific equations using TOBEC indices predicted lean and lipid masses of our sample of birds with average errors of 8-28% and 53-155%, respectively. A new two-species equation relating lean mass and TOBEC indices revealed average errors of 4.6% and 23.2% in predicting lean and lipid mass, respectively. New intraspecific equations that estimate lipid mass directly from body mass, morphological measurements, and TOBEC indices yielded about a 13% error in lipid estimates. Body mass and morphological measurements explained a substantial portion of the variance (about 90%) in fat mass of both species. Addition of TOBEC indices improved the predictive model more for the smaller than for the larger sandpiper. TOBEC indices explained an additional 7.8% and 2.6% of the variance in fat mass and reduced the minimum breadth of prediction intervals by 0.95 g (32%) and 0.39 g (13%) for Semipalmated and White-rumped Sandpipers, respectively. The breadth of prediction intervals for models used to predict fat levels of individual birds must be considered when interpreting the resultant lipid estimates.

Hatching asynchrony in American Goldfinches: an experimental study

I examined Lacks (1954, 1968) hypothesis that asynchronous hatching is an adaptive response to food shortage during the breeding season by comparing growth and survival of nestlings in asynchronous and artificially created synchronous broods of American Goldfinches (Carduelis tristis). I also examined the effects of seasonal and environmental factors on nestling growth and survival and on hatching asynchrony. Nestlings in asynchronous broods were more likely to diverge in mass and did so at a faster rate than those in synchronous broods. The lightest nestlings of asynchronous broods grew more slowly than their heavier nestmates and than all nestlings in synchronous broods. Brood reduction occurred more frequently in asynchronous broods. Survival and growth rates decreased throughout the breeding season and during inclement weather. Hatching intervals increased during the season. When brood reduction or differential growth among nestmates occurred in asynchronous broods, suggesting that the energy available for growth was limiting, heavy nestlings in asynchronous broods grew nonsignificantly faster than heavy nestlings in concurrent synchronous broods. This trend implies that when insufficient food is delivered to nestlings, asynchronous hatching may provide a slight advantage for older nestlings. Growth rates of all nestlings, however, were greatest in highly synchronous broods. Explanations other than growth of young must be invoked to explain why extreme synchrony is not more common in goldfinches. I discuss constraints on the normal pattern of hatching asynchrony characteristic of this species. The proximate mechanism for differential feeding and brood reduction is discussed.

LANDBIRD MIGRATION IN THE AMERICAN WEST: RECENT PROGRESS AND FUTURE RESEARCH DIRECTIONS

Abstract. Our knowledge of avian behaviors during the nonbreeding period still lags behind that of the breeding season, but the last decade has witnessed a proliferation in research that has yielded significant progress in understanding migration patterns of North American birds. And, although historically the great majority of migration research has been conducted in the eastern half of the continent, there has been much recent progress on aspects of avian migration in the West. In particular, expanded use of techniques such as radar, plasma metabolites, mist-netting, count surveys, stable isotopes, genetic data, and animal tracking, coupled with an increase in multi-investigator collaborations, have all contributed to this growth of knowledge. There is increasing recognition that migration is likely the most limiting time of year for migratory birds, increasing the importance of continuing to decipher patterns of stopover ecology, identifying critical stopover habitats, and documenting migration routes in the diverse and changing landscapes of the American West. Here, we review and briefly synthesize the latest findings and advances in avian migration and consider research needs to guide future research on migration in the West.

Reducing sedimentation of depressional wetlands in agricultural landscapes

Depressional wetlands in agricultural landscapes are easily degraded by sediments and contaminants accumulated from their watersheds. Several best management practices can reduce transport of sediments into wetlands, including the establishment of vegetative buffers. We summarize the sources, transport dynamics, and effect of sediments, nutrients, and contaminants that threaten wetlands and the current knowledge of design and usefulness of grass buffers for protecting isolated wetlands. Buffer effectiveness is dependent on several factors, including vegetation structure, buffer width, attributes of the surrounding watershed (i.e., area, vegetative cover, slope and topography, soil type and structure, soil moisture, amount of herbicides and pesticides applied), and intensity and duration of rain events. To reduce dissolved contaminants from runoff, the water must infiltrate the soil where microbes or other processes can break down or sequester contaminants. But increasing infiltration also diminishes total water volume entering a wetland, which presents threats to wetland hydrology in semi-arid regions. Buffer effectiveness may be enhanced significantly by implementing other best management practices (e.g., conservation tillage, balancing input with nutrient requirements for livestock and crops, precision application of chemicals) in the surrounding watershed to diminish soil erosion and associated contaminant runoff. Buffers require regular maintenance to remove sediment build-up and replace damaged or over-mature vegetation. Further research is needed to establish guidelines for effective buffer width and structure, and such efforts should entail a coordinated, regional, multi-scale, multidisciplinary approach to evaluate buffer effectiveness and impacts. Direct measures in “real-world” systems and field validations of buffer-effectiveness models are crucial next steps in evaluating how grass buffers will impact the abiotic and biotic variables attributes that characterize small, isolated wetlands.